Cited by 97Evolutionary Psychology and Human BehaviorLanguage and cultural evolutionAnimal Behavior and Reproduction

Hypotheses for the Evolution of Reduced Reactive Aggression in the Context of Human Self-Domestication

Open access
Wrangham, Richard W.·2019·Frontiers in Psychology

Abstract

Parallels in anatomy between humans and domesticated mammals suggest that for the last 300,000 years, Homo sapiens has experienced more intense selection against the propensity for reactive aggression than any other species of Homo. Selection against reactive aggression, a process that can also be called self-domestication, would help explain various physiological, behavioral and cognitive features of humans, including the unique system of egalitarian male hierarchy in mobile hunter-gatherers. Here I review nine leading proposals that could potentially explain why self-domestication occurred in H. sapiens, whether by selecting in favor of cooperation and tolerance or by selecting against aggressiveness. To account for the domestication syndrome, proposals must explain what led to a decline in fitness of highly aggressive males, and why the explanatory factor applies only to H. sapiens rather than to other species of Homo. The proposed explanations invoke genetic group selection; group-structured culture selection (also known as cultural group selection); social selection by female mate choice; social selection by male partner choice; increased self-control; cooperative breeding; high population density; use of lethal weapons; and language-based conspiracy. Most proposals face difficulties in accounting for the origins and/or maintenance of self-domestication. I conclude that the evolution of language-based conspiracy, which is a form of collective intentionality, was the key factor initiating and maintaining self-domestication in H. sapiens, because it is the most convincing mechanism for explaining the selective pressure against individually powerful fighters. Sophisticated language enabled males of low fighting prowess to coordinate in executing physically aggressive and domineering alpha males. This system is known today as a leveling mechanism in small-scale societies. Group-structured culture selection possibly accelerated the process.

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Written from the paper’s full text.

TL;DR

Wrangham reviews nine hypotheses for why *Homo sapiens* underwent self-domestication via reduced reactive aggression, arguing each must explain why effective alpha fighters failed to monopolize fitness and why this applied only to *H. sapiens*. He concludes that language-based conspiracy—coordinated planned execution of domineering males—best accounts for initiating and maintaining selection against reactive aggression, with group-structured culture selection possibly accelerating it.

Key findings

  • Anatomical evidence indicates a domestication syndrome in *H. sapiens*; the Jebel Irhoud specimens from around 315,000 years ago show short faces, small teeth, and reduced brow-ridges, suggesting selection for self-domestication began before 300,000 years ago. Within-group aggressive conflict frequencies in *H. sapiens* are two to three orders of magnitude lower than in wild chimpanzees and bonobos.
  • Of nine proposals reviewed, most face difficulties in accounting for the origins and/or maintenance of reduced reactive aggression: genetic group selection, group-structured culture selection, female mate choice, male partner choice, self-control, cooperative breeding, population density, and lethal weapons.
  • Language-based conspiracy is argued to be the key factor: sophisticated language enabled males of low fighting prowess to cooperatively plan the execution of physically aggressive and domineering alpha males, a system known today as a leveling mechanism in small-scale societies.
  • Group-structured culture selection possibly accelerated the process, but the paper concludes it is unlikely to have been important 300,000 years ago and does not by itself explain selection against aggression.

Methods

  • Conceptual review and hypothesis paper; no new empirical data are presented. It reviews nine leading proposals for self-domestication in *H. sapiens*, selected as prominent explanations for reduced aggressiveness or increased cooperation.
  • Each proposal is evaluated against two challenges: it must account for the failure of the most individually effective fighters to dominate access to resources and maximize fitness, and it must apply only to *H. sapiens*, since only this species is known to exhibit a domestication syndrome.
  • The analysis draws on comparative primate, fossil, anatomical, ethnographic, and experimental literature; Table 1 summarizes merits and problems for each scenario.

Limitations

  • Stated: the paper does not review any explanation in detail; the timing of language skills is speculative; the hypothesis is hard to test because relevant cultural practices are extinct; the ethnographic record of capital punishment is thin; and the composition of domestication syndromes is unpredictable, with uncertain biological mechanisms.
  • Stated: the conclusion that *H. sapiens* acquired low reactive aggression during the last 300,000 years can still be challenged on the basis that the supposed domestication syndrome may reflect independent adaptations or an earlier genus-wide adaptation.
  • Evident: as a review/hypothesis synthesis, it offers no primary quantitative test of execution rates or genetic selection, and it relies on present-day hunter-gatherers and nonhuman primates as analogs for past behavior.

Model-written summary. Check it against the paper before relying on it.

Integrity signals

Registry

Crossref · 10.3389/fpsyg.2019.01914

Cites

87 of 87 references verified0 citation errors · 0 not checkable

Statistics

Nothing to recompute.

Claims the paper makes

0 of 103 cited sources checked · 0 supported · 0 flagged

  • Interpretationreasoning

    Parallels in anatomy between humans and domesticated mammals suggest that for the last 300,000 years, Homo sapiens has experienced more intense selection against the propensity for reactive aggression than other species of Homo.

    “Parallels in anatomy between humans and domesticated mammals suggest that for the last 300,000 years, Homo sapiens has experienced more intense selection against the propensity for reactive aggression than other species of Homo.”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Leach, 2003 source

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    • cites Hare, 2017

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    • cites Wrangham, 2019

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  • Premisecitation

    The typical primate dominance system (alpha male achieving position by defeating lower-ranking males, dominating females, higher fitness) is found in chimpanzees, gorillas, savanna baboons, and most other primates living in multi-female, multi-male groups.

    “This system is found in chimpanzees (Wroblewski et al., 2009), gorillas (Gorilla spp.) (Nsubuga et al., 2008; Breuer et al., 2012), savanna baboons (Papio spp.) (Alberts et al., 2006; Baniel et al., 2017), and most other primates living in multi-female, multi-male groups.”
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    • cites Wroblewski et al., 2009 source

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    • cites Nsubuga et al., 2008 source

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  • Interpretationreasoning

    Pleistocene Homo experienced selection for self-domestication starting before 300,000 years ago.

    “These points suggest that Pleistocene Homo experienced selection for self-domestication starting before 300,000 years ago.”
    Quote in the paperSubstantial claim
  • Premisecitation

    All four features used by archeologists to recognize a domesticated species in the fossil record are found in H. sapiens.

    “Leach showed that all four traits are found in H. sapiens, although cranial capacity was not reduced until the end of the Pleistocene and the significance of its reduction has been challenged (Ruff et al., 1997).”
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    • cites Leach, 2003 source

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  • Interpretationreasoning

    The presence of a domestication syndrome appears to be a signal of selection against reactive aggression.

    “The presence of a domestication syndrome therefore appears to be a signal of selection again reactive aggression.”
    Quote in the paperSubstantial claim
  • Classified as probably not a substantial claim. 57
  • May combine two claims. 51
  • Probably not a substantial claim: may not pass S2 (checkable), S4 (based).

    Interpretationnone given

    Language-based conspiracy is the most convincing mechanism for explaining the selective pressure against individually powerful fighters.

    “because it is the most convincing mechanism for explaining the selective pressure against individually powerful fighters.”
    Quote in the paperNot a substantial claim
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    Premisecitation

    Facial width and brow-ridge projection have declined in H. sapiens.

    “Facial width and brow-ridge projection have also declined in H. sapiens (Cieri et al., 2014).”
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    • cites Cieri et al., 2014 source

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    Interpretationreasoning

    Group-structured culture selection possibly accelerated the process of self-domestication.

    “Group-structured culture selection possibly accelerated the process.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Premisecitation

    The Jebel Irhoud specimens have short faces, small teeth, and reduced brow-ridges compared to pre-sapiens ancestors, making them the earliest human specimens to show features of the domestication syndrome.

    “The Jebel Irhoud specimens have short faces, small teeth, and reduced brow-ridges compared to pre-sapiens ancestors, making them the earliest human specimens to show features of the domestication syndrome.”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Hublin et al., 2017 source

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    Interpretationreasoning

    Selection has acted to reduce the propensity for reactive aggression in bonobos compared to a chimpanzee-like ancestor, such that male bonobos who use reactive aggression to achieve an alpha position by being the most effective fighter would not tend to achieve maximal fitness.

    “In other words selection has acted to reduce the propensity for reactive aggression in bonobos, compared to a chimpanzee-like ancestor, such that male bonobos who use reactive aggression to achieve an alpha position by being the most effective fighter would not tend to achieve maximal fitness.”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The ancestral Homo species sometimes called Homo heidelbergensis lived in social communities and had dominance hierarchies typical of primates living in multi-female, multi-male groups.
    • The authors state this as an assumption: In the ancestral Homo species, there was an alpha male who achieved his position by physically defeating lower-ranking males in one-on-one combat.
    • The authors state this as an assumption: The alpha male dominated all females and experienced higher fitness than other males.
  • Classified as probably not a substantial claim.

    Premisecitation

    Features of the domestication syndrome were produced by selection purely for docility, i.e., for a low propensity for reactive aggression.

    “Belyaev (1969) showed that features of the domestication syndrome were produced by selection purely for docility, i.e., for a low propensity for reactive aggression (Trut, 1999).”
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    Premisecitation

    Marean (2015) developed a version of the group selection hypothesis suggesting why selection in favor of cooperation was more intense in the origin of H. sapiens than in other Homo, pointing to increasing exploitation of dense and predictable food resources, leading to higher population density, and more aggressive defense of territories.

    “Marean (2015) developed a version of this hypothesis that suggested why selection in favor of cooperation was more intense in the origin of H. sapiens than in other Homo. He pointed to increasing exploitation of dense and predictable food resources, leading to higher population density, and more aggressive defense of territories.”
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    • cites Marean, 2015 source

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  • Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based). May combine two claims.

    Interpretationnone given

    The explanation for selection against reactive aggression must apply only to H. sapiens, since it is the only species of Homo known to exhibit a domestication syndrome.

    “The second challenge is that the explanation for selection against reactive aggression must apply only to H. sapiens, since it is the only species of Homo known to exhibit a domestication syndrome.”
    Quote in the paperNot a substantial claim
  • Classified as probably not a substantial claim.

    Interpretationreasoning

    If the male-coalition suppression mechanism applied throughout the existence of H. sapiens, it would explain how there was selection against the best individual fighters.

    “If this mechanism applied throughout the existence of H. sapiens, it would explain how there was selection against the best individual fighters [see (8) and (9) below].”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: The present informs the past.
    • Rests on an assumption: Contemporary mobile hunter-gatherer societies offer a guide to reconstructing late Pleistocene societies.
  • May combine two claims.

    Premisecitation

    Living adult males with relatively narrower faces tend both to be less reactively aggressive and to be perceived as being less aggressive, to the point of being assessed as having more human-like minds.

    “In support of the behavioral significance of recent anatomical changes, living adult males with relatively narrower faces tend both to be less reactively aggressive and to be perceived as being less aggressive (Carré and McCormick, 2008; Haselhuhn et al., 2015), to the point of being assessed as having more human-like minds (Deska et al., 2018).”
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    • cites Carré and McCormick, 2008

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    • cites Haselhuhn et al., 2015 source

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    Premisecitation

    The high degree of docility characteristic of humans and domesticated animals depends on a low propensity for reactive aggression.

    “The high degree of docility that is characteristic of humans and domesticated animals depends on a low propensity for reactive aggression, but what relationship it has with proactive aggression, if any, is unknown (Wrangham, 2018).”
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    • cites Wrangham, 2018 source

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    Interpretationnone given

    There is reason to question the core assumption that success in intergroup conflict is increased by a reduction in within-group conflict.

    “Finally, there is reason to question the core assumption that success in intergroup conflict is increased by a reduction in within-group conflict.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Interpretationreasoning

    In recent cases GSCS has little relevance for genetic evolution of populations, and none for genetic evolution of the species as a whole.

    “In those cases it has little relevance for genetic evolution of populations, and none for genetic evolution of the species as a whole.”
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  • Classified as probably not a substantial claim.

    Interpretationreasoning

    Hypotheses for the evolution of increased cooperation could in theory contribute to explaining why docility evolved.

    “Hypotheses for the evolution of increased cooperation could therefore in theory contribute to explaining why docility evolved (e.g., Henrich, 2016; Richerson et al., 2016).”
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    • cites Henrich, 2016 source

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    • cites Richerson et al., 2016 source

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    Interpretationreasoning

    Sophisticated language enabled males of low fighting prowess to cooperatively plan the execution of physically aggressive and domineering alpha males.

    “Sophisticated language enabled males of low fighting prowess to cooperatively plan the execution of physically aggressive and domineering alpha males.”
    Quote in the paperNot a substantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    The earliest record of a lineage of H. sapiens comes from around 315,000 years ago from Jebel Irhoud, Morocco.

    “Currently the earliest record of a lineage of H. sapiens comes from around 315,000 years ago from Jebel Irhoud, Morocco (Hublin et al., 2017).”
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    • cites Hublin et al., 2017 source

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    Premisecitation

    In bonobos, the alpha male ordinarily owes his position to the support of his mother rather than to his ability to fight alone, and he is not necessarily dominant to the alpha female.

    “In bonobos, there is an alpha male, but he ordinarily owes his position to the support of his mother rather than to his ability to fight alone, and he is not necessarily dominant to the alpha female (Surbeck et al., 2011).”
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    • cites Surbeck et al., 2011 source

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    Premisecitation

    In some human societies a coalition of the majority prevents any individuals from dominating access to preferred resources, and there are no alpha males (reverse dominance hierarchy).

    “In some societies a coalition of the majority is able to prevent any individuals from dominating access to preferred resources (Boehm, 1999). In these cases, called a “reverse dominancy hierarchy” or “counter-hierarchy” there are no alpha males (Boehm, 1993).”
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    • cites Boehm, 1999 source

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    • cites Boehm, 1993 source

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    Interpretationreasoning

    The group selection argument does not differentiate between humans and chimpanzees, even though only humans, and not chimpanzees, have experienced selection against reactive aggression.

    “The group selection argument thus does not differentiate between humans and chimpanzees, even though only humans, and not chimpanzees, have experienced selection against reactive aggression.”
    Quote in the paperSubstantial claim
  • Classified as probably not a substantial claim. May combine two claims.

    Premisecitation

    Human aggressiveness comes in two major forms, reactive (or impulsive) and proactive (or premeditated), each with their own distinctive neurobiology.

    “Human aggressiveness has recently been argued to come in two major forms, reactive (or impulsive) and proactive (or premeditated), each with their own distinctive neurobiology (Wrangham, 2018).”
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    • cites Wrangham, 2018 source

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    Premisecitation

    Group selection has been proposed as a mechanism for promoting parochial altruism.

    “Group selection has been proposed as a mechanism for promoting parochial altruism, meaning altruism restricted to “one’s own ethnic, racial, or other group” (Choi and Bowles, 2007: 636).”
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    Interpretationnone given

    There is no direct evidence as to whether, around the time that H. sapiens evolved, foods could no longer be collected individually, teamwork against predators became much more important, or pair-bonding began.

    “For example there is no direct evidence as to whether, around the time that H. sapiens evolved, foods could no longer be collected individually, teamwork against predators became much more important, or pair-bonding began.”
    Quote in the paperNot a substantial claim
  • Probably not a substantial claim: may not pass S1 (asserted).

    Premisecitation

    The argument is that cooperative breeding was a feature of the genus Homo from H. erectus onward, close to 2 million years ago.

    “First, the argument made by Hrdy (2009), Burkart et al. (2009), and Burkart and van Schaik (2016) is that cooperative breeding was a feature of the genus Homo from H. erectus onward, close to 2 million years ago.”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites Hrdy, 2009

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    • cites Burkart et al., 2009 source

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    • cites Burkart and van Schaik, 2016

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    Interpretationreasoning

    Hypotheses of social selection for unaggressive partners are challenged by the predictable success of a tyrant using brute force for personal gain.

    “In short, hypotheses of social selection for unaggressive partners are challenged by the predictable success of a tyrant using brute force for personal gain.”
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    Premisecitation

    Marean developed a similar scenario.

    “Marean (2015) developed a similar scenario.”
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    • cites Marean, 2015 source

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    Premisecitation

    Selection favored a low propensity for male aggression because offspring were thereby better protected from the father's potential violence.

    “Selection favored a low propensity for male aggression because offspring were thereby better protected from the father’s potential violence.”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Tomasello, 2016 source

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    Premisecitation

    In societies with higher-status women, men were relatively short, suggesting reduced selection for male aggressiveness.

    “In support, they found that in societies with higher-status women (who were presumed to have more freedom to choose their mates), men were relatively short (suggesting reduced selection for male aggressiveness).”
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    • cites Gleeson and Kushnick, 2018

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    Premisecitation

    Increasing population density around 200,000 years ago could have been responsible for selection pressures against reactive aggression.

    “Cieri et al. (2014) proposed that increasing population density around 200,000 years ago could have been responsible for selection pressures against reactive aggression.”
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    • cites Cieri et al., 2014 source

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  • Probably not a substantial claim: may not pass S4 (based).

    Interpretationreasoning

    If group cultures in the form of social norms occurred as early as 300,000 years ago, GSCS might have contributed to selection against reactive aggression.

    “Nevertheless, if group cultures in the form of social norms occurred as early as 300,000 years ago, GSCS might have contributed to selection against reactive aggression.”
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  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Increasing population density was plausibly associated with the cultural developments around 300,000 years ago.

    “In favor of this general kind of idea, increasing population density was plausibly associated with the cultural developments reported by Brooks et al. (2018) around 300,000 years ago.”
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    • cites Brooks et al., 2018 source

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    Interpretationreasoning

    The self-control hypothesis is challenged by the problems of why the sapiens lineage took a different course from other Homo, and by the problem of how increased self-control would lead to selection against domineering behaviors.

    “It is challenged by the problems of why the sapiens lineage took a different course from other Homo, and by the problem of how increased self-control would lead to selection against domineering behaviors.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S1 (asserted). May combine two claims.

    Premisecitation

    Higher population density could have increased the pressure to exploit resources controlled by other humans, and would have selected for increased social tolerance toward unfamiliar individuals, an extension of social networks, and more sharing of acquired foods.

    “They suggested that because higher population density could have increased the pressure to exploit resources controlled by other humans, it would have selected for increased social tolerance toward unfamiliar individuals, an extension of social networks, and more sharing of acquired foods.”
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    • cites Cieri et al., 2014 source

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    Premisereasoning

    Effective cooperation depended on a reduction in selfishness and aggressiveness, and on a concomitant increase in sympathy and shared intentionality.

    “Effective cooperation depended on a reduction in selfishness and aggressiveness, and on a concomitant increase in sympathy and shared intentionality.”
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    • cites Tomasello, 2016 source

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  • Probably not a substantial claim: may not pass S1 (asserted), S2 (checkable), S3 (load-bearing), S4 (based). May combine two claims.

    Interpretationnone given

    An alternative proposal that could have involved GSCS attributes selection against reactive aggression to the emergence of language-based conspiracy that would allow execution of aggressive individuals.

    “An alternative proposal that could have involved GSCS attributes selection against reactive aggression to the emergence of language-based conspiracy that would allow execution of aggressive individuals.”
    Quote in the paperNot a substantial claim
  • Probably not a substantial claim: may not pass S1 (asserted).

    Premisecitation

    Self-domestication could be promoted by an increased ability for self-control leading to less use of reactive aggression.

    “Based on the latter finding, Hare (2017) suggested that self-domestication could be promoted by an increased ability for self-control leading to less use of reactive aggression.”
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    • cites Hare, 2017

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  • Probably not a substantial claim: may not pass S4 (based).

    Interpretationnone given

    Tomasello does not discuss what would stop a determined, physically powerful male from exerting his fighting ability at the expense of others in his group.

    “Similar to the problem with the female mate choice hypothesis, the difficulty is that Tomasello does not discuss what would stop a determined, physically powerful male from exerting his fighting ability at the expense of others in his group.”
    Quote in the paperNot a substantial claim
  • Classified as probably not a substantial claim. May combine two claims.

    Interpretationreasoning

    Applying the hypothesis specifically to self-domestication, selected effects could in theory include reduced tendencies for reactive aggression, on the traditional premise that groups in which males competed more aggressively with group members would be less effective at intergroup conflict.

    “Applying the hypothesis specifically to the problem of self-domestication, selected effects could in theory include reduced tendencies for reactive aggression, on the traditional premise that groups in which males competed more aggressively with group members would be less effective at intergroup conflict (e.g., Zhang et al., 2019).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Zhang et al., 2019 source

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  • May combine two claims.

    Interpretationreasoning

    The cooperative breeding proposal has no special claim on events surrounding the evolution of H. sapiens around 300,000 years ago: it does not help explain the evidence that H. sapiens underwent a singular form of self-domestication not experienced by other Homo species.

    “Their proposal therefore has no special claim on events surrounding the evolution of H. sapiens around 300,000 years ago: it does not help explain the evidence that H. sapiens underwent a singular form of self-domestication not experienced by other Homo species.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    In experimental studies with captive primates, population density is sometimes associated with reduced aggression and increased tolerance.

    “In experimental studies with captive primates, population density is sometimes associated with reduced aggression and increased tolerance (Aureli et al., 2000).”
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    • cites Aureli et al., 2000 source

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  • Probably not a substantial claim: may not pass S2 (checkable).

    Interpretationreasoning

    Such findings mean that a specific theory is needed to explain why intergroup conflict would have led to reactive aggression being selected against more intensely in Homo ∼300,000 years ago than happens in other primates.

    “Such findings mean that a specific theory is needed to explain why intergroup conflict would have led to reactive aggression being selected against more intensely in Homo,∼300,000 years ago, than happens in other primates.”
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    Premisecitation

    Tomasello presented an explanation for the evolution of morality that included a proposal for the reduction of aggressiveness in H. sapiens, called the interdependence hypothesis.

    “This general idea has been elaborated particularly clearly by Tomasello (2016), who presented an explanation for the evolution of morality that included a proposal for the reduction of aggressiveness in H. sapiens. Tomasello called it the “interdependence hypothesis.””
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    • cites Tomasello, 2016 source

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  • Classified as probably not a substantial claim. May combine two claims.

    Premisecitation

    In primates, absolute brain size predicts neuron number and is strongly correlated with the ability to inhibit prepotent responses.

    “In primates, absolute brain size predicts neuron number and is strongly correlated with the ability to inhibit prepotent responses (MacLean et al., 2014).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites MacLean et al., 2014 source

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    Premisecitation

    Gene-culture coevolution would then include selection in favor of an increasingly parochial-altruistic psychology: tendencies for docility and conformity would be favored because individuals with those characteristics would readily acquire group norms.

    “Gene-culture coevolution would then include selection in favor of an increasingly parochial-altruistic psychology: tendencies for docility and conformity would be favored because individuals with those characteristics would readily acquire group norms (Richerson et al., 2016).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Richerson et al., 2016 source

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  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Premisecitation

    Female choice of less aggressive males as mates was proposed as a mechanism that could promote self-domestication, because females who choose less aggressive males can benefit by their mates' greater investment in shared parenting effort.

    “Female choice of less aggressive males as mates was proposed by Cieri et al. (2014) and Gleeson and Kushnick (2018) as a mechanism that could promote self-domestication, because females who choose less aggressive males can benefit by their mates’ greater investment in shared parenting effort.”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Cieri et al., 2014 source

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    • cites Gleeson and Kushnick, 2018

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    Premisecitation

    Island populations of rodents, lizards and birds tend to live at higher density than their continental relatives and also show low levels of aggression.

    “Island populations tend to live at higher density than their continental relatives, and also to show low levels of aggression (Stamps and Buechner, 1985).”
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    • cites Stamps and Buechner, 1985

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    Premisecitation

    According to the interdependence hypothesis, the initial influence on the road to a human style of morality occurred around 400,000 years ago when there was a disappearance of individually obtainable foods.

    “According to the interdependence hypothesis the initial influence on the road to a human style of morality occurred around 400,000 years ago when there was a “disappearance of individually obtainable foods” (Tomasello, 2016: 136).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Tomasello, 2016 source

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based).

    Interpretationreasoning

    Like a male lion feeding off the kills brought down by females, the determinedly aggressive and effective fighter would continue to have high fitness thanks to his ability to commandeer food, mates or other resources from others in the group.

    “Like a male lion Panthera leo feeding off the kills brought down by females, the determinedly aggressive and effective fighter would continue to have high fitness thanks to his ability to commandeer food, mates or other resources from others in the group.”
    Quote in the paperNot a substantial claim
  • Probably not a substantial claim: may not pass S4 (based). May combine two claims.

    Interpretationreasoning

    The despotic male would improve his lot by seizing the choicest foods that others produced: his physical dominance would allow it.

    “The despotic male would improve his lot by seizing the choicest foods that others produced: his physical dominance would allow it.”
    Quote in the paperNot a substantial claim
  • Probably not a substantial claim: may not pass S4 (based).

    Interpretationreasoning

    The only obvious solution would be that the tyrant is constrained by coalitions that can physically rein in his power.

    “The only obvious solution would be that the tyrant is constrained by coalitions that can physically rein in his power, similar to the solution for social selection by female mate choice.”
    Quote in the paperNot a substantial claim
  • Classified as probably not a substantial claim.

    Premisecitation

    The tribal social instincts hypothesis suggests that groups in which there were norms of parochial altruism would have been especially successful in intergroup competition.

    “This hypothesis suggests that groups in which there were norms of parochial altruism would have been especially successful in intergroup competition.”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Richerson and Boyd, 1998

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S4 (based).

    Interpretationnone given

    Increased social tolerance implies a reduced propensity for reactive aggression.

    “Increased social tolerance implies a reduced propensity for reactive aggression.”
    Quote in the paperNot a substantial claim
  • Probably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing).

    Interpretationreasoning

    The notion of a relevant ecological change is entirely speculative.

    “Thus the notion of a relevant ecological change is entirely speculative.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Premisecitation

    Effective population size of H. sapiens fell from 200,000 years ago to 50,000-30,000 years ago, suggesting a mismatch between cultural development and population density.

    “On the other hand, genetic data indicate that effective population size of H. sapiens fell from 200,000 years ago to 50,000–30,000 years ago, suggesting a mismatch between cultural development and population density (Steele and Weaver, 2014).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Steele and Weaver, 2014

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based). May combine two claims.

    Interpretationnone given

    While such selection is theoretically possible, no cases have yet been documented of GSCS leading to genetic change.

    “While such selection is theoretically possible, to my knowledge no cases have yet been documented of GSCS leading to genetic change.”
    Quote in the paperNot a substantial claim
  • Classified as probably not a substantial claim. May combine two claims.

    Premisecitation

    Cooperative breeding became an important feature of early Homo society and selected for increased social tolerance.

    “Burkart et al. (2009), Hrdy (2009), and Burkart and van Schaik (2016) proposed that cooperative breeding, a social system in which individuals help rear others’ offspring at a cost to their own reproductive effort, became an important feature of early Homo society and selected for increased social tolerance.”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Burkart et al., 2009 source

      Not checked in the free run.

    • cites Hrdy, 2009

      Not checked in the free run.

    • cites Burkart and van Schaik, 2016

      Not checked in the free run.

  • May combine two claims.

    Interpretationreasoning

    Coalitionary support is needed to enable partner choice to be effective, but coalitionary support has not featured to date in the social selection hypotheses.

    “Coalitionary support is needed to enable partner choice to be effective, but coalitionary support has not featured to date in the social selection hypotheses.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Interpretationreasoning

    Language-based conspiracy appears to be a vital prerequisite for controlling a would-be despot in a mobile hunter-gatherer society, and seems likely to have been equally important in the past.

    “For those reasons, language-based conspiracy appears to be a vital prerequisite for controlling a would-be despot in a mobile hunter-gatherer society, and seems likely to have been equally important in the past.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Premisecitation

    In humans, groups conspire against resented tyrants; in mobile hunter-gatherers plans are made and tyrants are executed.

    “In humans, by contrast, groups conspire against resented tyrants: in mobile hunter-gatherers plans are made and tyrants are executed.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based).

    Interpretationnone given

    Lethal weaponry would have facilitated executions.

    “Lethal weaponry would also have facilitated executions.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Premisecitation

    Experimental studies of captive primates show that high population density can lead to an increase in the rate of aggression, or can have no effect.

    “and experimental studies of captive primates show that high population density can lead to an increase in the rate of aggression, or can have no effect (Cordoni and Palagi, 2007; Crast et al., 2015).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Cordoni and Palagi, 2007

      Not checked in the free run.

    • cites Crast et al., 2015 source

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Interpretationreasoning

    The lethal weapons hypothesis has the merit of suggesting that the use of lethal weapons would lead to selection against reactive aggression.

    “Since alpha-males would have maintained their position by reacting violently to challengers, this hypothesis has the merit of suggesting that the use of lethal weapons would lead to selection against reactive aggression.”
    Quote in the paperNot a substantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based).

    Interpretationnone given

    The time when language became sufficiently sophisticated that subordinate males could confidently plan to kill a better fighter is speculative.

    “As for language-based conspiracy (9), the time when language became sufficiently sophisticated that subordinate males could confidently plan to kill a better fighter is speculative.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Interpretationreasoning

    In groups that have egalitarian male dominance hierarchies, individual choice of unaggressive mates and unaggressive task partners are plausible influences on the reduction of reactive aggression.

    “In groups that have egalitarian male dominance hierarchies both ideas are plausible influences on the reduction of reactive aggression.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Oldowan stone manuports were possibly lethal throwing weapons in the Lower Pleistocene.

    “Oldowan stone manuports were possibly lethal throwing weapons in the Lower Pleistocene (Gintis et al., 2015).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Gintis et al., 2015 source

      Not checked in the free run.

  • May combine two claims.

    Interpretationreasoning

    The evolution of this newly sophisticated cognitive ability would have led subordinates to socially select against aggressive fighters, creating a reverse dominance hierarchy.

    “The evolution of this newly sophisticated cognitive ability would have led subordinates to socially select against aggressive fighters, creating a reverse dominance hierarchy.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Premisecitation

    Across species, the precise composition of domestication syndromes is unpredictable, undermining confidence in using them to infer selection against reactive aggression; and the biological mechanisms responsible for producing domestication syndromes are uncertain.

    “Against this idea, across species the precise composition of domestication syndromes is unpredictable, a fact that undermines confidence in using them to infer selection against reactive aggression; and the biological mechanisms responsible for producing domestication syndromes are uncertain (Sánchez-Villagra et al., 2016; Sánchez-Villagra and van Schaik, 2019).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Sánchez-Villagra et al., 2016 source

      Not checked in the free run.

    • cites Sánchez-Villagra and van Schaik, 2019 source

      Not checked in the free run.

  • May combine two claims.

    Interpretationreasoning

    Language-based conspiracy is not present in chimpanzees: subordinate chimpanzees cannot plan premeditated take-downs of alpha males, which probably partly explains why there has apparently been no selection against reactive aggression in this species.

    “Language-based conspiracy is not present in chimpanzees: subordinate chimpanzees cannot plan premeditated take-downs of alpha males, which probably partly explains why there has apparently been no selection against reactive aggression in this species.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Interpretationcitation

    Once reactive aggression started to be selected against, expected effects would have included, as a by-product, increased social tolerance and improved abilities to communicate and cooperate.

    “Once reactive aggression started to be selected against, expected effects would have included, as a by-product, increased social tolerance and improved abilities to communicate and cooperate (Hare, 2017).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Hare, 2017

      Not checked in the free run.

  • Classified as probably not a substantial claim.

    Interpretationcitation

    Sufficient execution of Pleistocene despots would have led to selection against reactive aggression.

    “Sufficient execution of Pleistocene despots would have led to selection against reactive aggression (Wrangham, 2018, 2019).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Wrangham, 2018, 2019

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Interpretationreasoning

    Executions can be planned using proactive aggression, so executioners arrange to kill a victim in a circumstance in which fighting back is essentially impossible, and killers incur very low costs.

    “An important feature of executions is that they can be planned using proactive aggression. This means that the executioners arrange to kill a victim in a circumstance in which fighting back is essentially impossible. As a result, the killers incur very low costs.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Interpretationreasoning

    The language-based conspiracy hypothesis can explain why reactive aggression would have been selected against, why this occurred only in H. sapiens, and why reactive aggression has continued to decline.

    “In sum, the language-based conspiracy hypothesis can explain why reactive aggression would have been selected against, why this occurred only in H. sapiens, and (given that capital punishment was recorded into contemporary times among mobile hunter-gatherers) why reactive aggression has continued to decline.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S4 (based).

    Interpretationnone given

    With weapon use already in place, capital punishment would have been relatively easy.

    “With weapon use already in place, capital punishment would have been relatively easy.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Premisecitation

    No other selective pressures are known to produce a domestication syndrome, and the resemblances that humans show to domesticated animals include physiological, behavioral and cognitive traits in addition to reduced aggression and a derived anatomy.

    “Nevertheless, no other selective pressures are known to produce a domestication syndrome, and the resemblances that humans show to domesticated animals include physiological, behavioral and cognitive traits in addition to reduced aggression and a derived anatomy (Hare, 2017; Wrangham, 2019).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Hare, 2017

      Not checked in the free run.

    • cites Wrangham, 2019

      Not checked in the free run.

  • Classified as probably not a substantial claim.

    Interpretationnone given

    The population density hypothesis fails to detail any process responsible for selection directly against alpha-male-style behavior.

    “The population density hypothesis also fails to detail any process responsible for selection directly against alpha-male-style behavior.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Interpretationnone given

    Current evidence about the evolutionary history of human population density undermines any easy correlations, and the relationship between population density and rates of aggression in primates is inconsistent.

    “Current evidence about the evolutionary history of human population density, however, undermines any easy correlations, and the relationship between population density and rates of aggression in primates is inconsistent.”
    Quote in the paperNot a substantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Interpretationnone given

    The cooperative breeding hypothesis could in theory account for reduced aggressiveness, although the evidence for cooperative breeders being unaggressive is inconsistent at best.

    “The cooperative breeding hypothesis for human social evolution (6) could in theory account for reduced aggressiveness, although the evidence for cooperative breeders being unaggressive is inconsistent at best.”
    Quote in the paperNot a substantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Interpretationcitation

    Increasing population density is associated with reduced aggressiveness in island species, and could in theory play a role in H. sapiens' self-domestication.

    “Increasing population density (7) is associated with reduced aggressiveness in island species, and could in theory play a role in H. sapiens’ self-domestication.”
    Quote in the paperNot a substantial claim
  • Probably not a substantial claim: may not pass S2 (checkable). May combine two claims.

    Interpretationreasoning

    The spread of the new style of hierarchy could have occurred by individual learning or by selection of group-cultures, and would have paved the way for diverse selection pressures to additionally influence the evolution of characteristically human social traits.

    “The spread of the new style of hierarchy could have occurred by individual learning or by selection of group-cultures, and would have paved the way for diverse selection pressures, as shown in Table 1, to additionally influence the evolution of the characteristically human social traits.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Premisecitation

    A biological hypothesis for the production of domestication syndromes has received important support, including evidence that genetic variants underlying self-domestication occur in H. sapiens but not in H. neanderthalensis or Denisovans.

    “Furthermore a biological hypothesis for the production of domestication syndromes (Wilkins et al., 2014) has received important support (Wilkins, 2017; Pendleton et al., 2018), including evidence that genetic variants underlying self-domestication occur in H. sapiens but not in H. neanderthalensis or Denisovans (Theofanopoulou et al., 2017).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Wilkins et al., 2014 source

      Not checked in the free run.

    • cites Wilkins, 2017 source

      Not checked in the free run.

    • cites Theofanopoulou et al., 2017 source

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    Capital punishment has been recorded among mobile hunter-gatherers in every continent, and aggressive bullies are a common type of victim.

    “This scenario is supported by the fact that capital punishment has been recorded among mobile hunter-gatherers in every continent, and that aggressive bullies are a common type of victim (Boehm, 2012).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Boehm, 2012

      Not checked in the free run.

  • May combine two claims.

    Interpretationreasoning

    Lethal weaponry does not appear to be associated with H. sapiens as tightly or as uniquely as theory would demand, since it has likely been used for a long time before the evolution of H. sapiens, as well as in other species of Homo.

    “However, it does not appear to be associated with H. sapiens as tightly or as uniquely as theory would demand, since it has likely been used for a long time before the evolution of H. sapiens, as well as in other species of Homo.”
    Quote in the paperNot a substantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based). May combine two claims.

    Interpretationnone given

    The use of lethal weapons was probably widespread from H. erectus onward, and would have facilitated the deliberate killing of conspecifics.

    “In sum, the use of lethal weapons was probably widespread from H. erectus onward, and would have facilitated the deliberate killing of conspecifics.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Interpretationreasoning

    Genetic group selection and group-structured culture selection do not account for reduced reactive aggression in H. sapiens in their current form, partly because in humans and other primates effective intergroup conflict is not necessarily associated with reduced within-group aggression.

    “I concluded that they do not do so in their current form, partly because in humans, and other primates, effective intergroup conflict is not necessarily associated with reduced within-group aggression.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S1 (asserted).

    Interpretationcitation

    Linguistic ability was significantly more sophisticated in H. sapiens than in H. neanderthalensis or other Homo species.

    “The prediction made by this hypothesis is that linguistic ability was significantly more sophisticated in H. sapiens than in H. neanderthalensis or other Homo species. That idea is plausible (Tattersall, 2016), though hotly debated (Dediu and Levinson, 2013).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Tattersall, 2016 source

      Not checked in the free run.

    • cites Dediu and Levinson, 2013

      Not checked in the free run.

  • May combine two claims.

    Interpretationcitation

    Because proactive and reactive aggression involve different neurobiological mechanisms, under a selective regime of alpha males being executed, the propensity for proactive reaction can remain high while the propensity for reactive aggression declines over evolutionary time.

    “The fact that proactive and reactive aggression involve different neurobiological mechanisms means that under a selective regime of alpha males being executed, the propensity for proactive reaction can remain high while the propensity for reactive aggression declines over evolutionary time (Wrangham, 2018).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Wrangham, 2018 source

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based). May combine two claims.

    Premisenone given

    The ethnographic record of capital punishment is thin; the practice has tended to disappear very rapidly.

    “Unfortunately, however, the ethnographic record of capital punishment is thin: the practice has tended to disappear very rapidly.”
    Quote in the paperNot a substantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based).

    Premisenone given

    Lethal weaponry has undoubtedly played an important role in human life.

    “Lethal weaponry (8) has undoubtedly played an important role in human life.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Interpretationreasoning

    The group selection argument for parochial altruism does not explain behavioral differences between humans and chimpanzees; and the culture selection argument for tribal social instincts is challenged by the origin of H. sapiens being so early that the requisite cultural abilities may not have been present.

    “The group selection argument for parochial altruism also does not explain behavioral differences between humans and chimpanzees; and the culture selection argument for tribal social instincts is challenged by the origin of H. sapiens being so early that the requisite cultural abilities may not have been present.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Premisecitation

    Chimpanzees cannot communicate to others that they wish to kill a particular individual, nor justify their desire, find out if their partner feels the same way, or plan to meet at a future time and place to carry out the deed.

    “Chimpanzees cannot communicate to others that they wish to kill a particular individual, let alone justify their desire, find out if their partner feels the same way, or plan to meet at some future time at a specified place in order to carry out the deed.”
    Quote in the paperSubstantial claim
89 of 89
  • The authors state this as an assumption.

    The ancestral Homo species sometimes called Homo heidelbergensis lived in social communities and had dominance hierarchies typical of primates living in multi-female, multi-male groups.

    “I follow the assumption (e.g., Gintis et al., 2015) that this ancestral Homo species, sometimes called Homo heidelbergensis, would have lived in social communities and would have had dominance hierarchies typical of primates living in multi-female, multi-male groups.”
  • The authors state this as an assumption.

    In the ancestral Homo species, there was an alpha male who achieved his position by physically defeating lower-ranking males in one-on-one combat.

    “Specifically there would have been an alpha male who achieved his position by physically defeating lower-ranking males in one-on-one combat.”
  • The authors state this as an assumption.

    The alpha male dominated all females and experienced higher fitness than other males.

    “The alpha would also have dominated all females, and would have predictably experienced higher fitness than other males.”
  • The authors state this as an assumption.

    The present informs the past.

    “The simplest hypothesis is based on the assumption that the present informs the past.”
  • The paper relies on this without support.

    Contemporary mobile hunter-gatherer societies offer a guide to reconstructing late Pleistocene societies.

    “Similarly, the societies of contemporary mobile hunter-gatherers offer a guide to reconstructing late Pleistocene societies.”
  • The authors state this as an assumption.

    A female mid-Pleistocene Homo would not have had the physical ability to freely express her mate choice.

    “Assuming that she would not have had the physical ability to freely express her choice, she would have had to rely on coalitionary help.”
  • The authors state this as an assumption.

    Humans benefit by cooperating in gathering, hunting, warfare and other tasks.

    “On the assumption that humans benefit by cooperating in gathering, hunting, warfare and other tasks, social selection has likely been important in promoting altruistic, and non-competitive behavior within groups (Nesse, 2009).”
  • The authors state this as an assumption.

    The anatomical signal of a domestication syndrome in H. sapiens is a reliable indicator of reduced reactive aggression.

    “The conceptual framework assumes that the anatomical signal of a domestication syndrome in H. sapiens is a reliable indicator of reduced reactive aggression.”
8 of 8

Claims other papers make about this one

Each is checked against this paper’s own text: does it say what the citing paper says it does?

  • Modern humans have markedly reduced intraspecific reactive aggression when compared to extant primates.

    “In conjunction with findings that our species has markedly reduced intraspecific reactive aggression when compared to extant primates, this has helped to spur research into the hypothesis that, relative to archaic hominins, modern humans have undergone positive selection for a reduction in reactive aggression towards each other [13].”

Cites

Flagged first, then by citations

Research works printed but not linked: 1

  1. Steele, 2014, Comment on Cieri et al. Craniofacial feminization, social tolerance, and the origins of behavioral modernity., Curr. Anthropol., 55, 434, 10.1086/677209 · DOI · Paper page

Cited by

Sorted by citations

Authors
Wrangham, R. W.
Affiliations
Harvard University (US)
Corresponding author
Wrangham, Richard W.
Venue
Frontiers in Psychology
Published
2019-08-20
Volume
10
Pages
1914-1914
ISSN
1664-1078 (ISSN-L)
OpenAlex
W2964997421
Cited by
97
References
87
Citations by year
2026: 10 · 2025: 16 · 2024: 14 · 2023: 14 · 2022: 17 · 2021: 16 · 2020: 8 · 2019: 1
FWCI
12.38
Type
journal article
Language
en
Open access
Locations
  • Frontiers in Psychology (publishedVersion, cc-by)
  • PubMed Central (publishedVersion, cc-by)
  • PubMed (publishedVersion)
  • DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion)
Funders
Kempe Foundation
Topics
  • Social Sciences › Psychology › Experimental and Cognitive Psychology › Evolutionary Psychology and Human Behavior
  • Social Sciences › Cultural Studies › Language and cultural evolution
  • Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Animal Behavior and Reproduction
Keywords
self-domestication, reactive aggression, human evolution, domestication syndrome, hunter-gatherers, group selection, cultural group selection, female mate choice, self-control, cooperative breeding, population density, lethal weapons, collective intentionality

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Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.

Claims read by deepseek-v4.1-flash under definition version 3 (extract-v3) on 2026-10-10.

Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-10-10.

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